Application of species richness estimators for the assessment of fungal diversity

Species richness and distribution patterns of wood-inhabiting fungi and mycetozoans (slime moulds) were investigated in the canopy of a Central European temperate mixed deciduous forest. Species richness was described with diversity indices and species-accumulation curves. Nonmetrical multidimensional scaling was used to assess fungal species composition on different tree species. Different species richness estimators were used to extrapolate species richness beyond our own data. The reliability of the abundance-based coverage estimator, Chao, Jackknife and other estimators of species richness was evaluated for mycological surveys. While the species-accumulation curve of mycetozoans came close to saturation, that of wood-inhabiting fungi was continuously rising. The Chao 2 richness estimator was considered most appropriate to predict the number of species at the investigation site if sampling were continued. Gray's predictor of species richness should be used if statements of the number of species in larger areas are required. Multivariate analysis revealed the importance of different tree species for the conservation and maintenance of fungal diversity within forests, because each tree species possessed a characteristic fungal community. The described mathematical approaches of estimating species richness possess great potential to address fungal diversity on a regional, national, and global scale.     

LiDAR GEDI derived tree canopy height heterogeneity reveals patterns of biodiversity in forest ecosystems

The “Height Variation Hypothesis” is an indirect approach used to estimate forest biodiversity through remote sensing data, stating that greater tree height heterogeneity (HH) measured by CHM LiDAR data indicates higher forest structure complexity and tree species diversity. This approach has traditionally been analyzed using only airborne LiDAR data, which limits its application to the availability of the dedicated flight campaigns. In this study we analyzed the relationship between tree species diversity and HH, calculated with four different heterogeneity indices using two freely available CHMs derived from the new space-borne GEDI LiDAR data. The first, with a spatial resolution of 30 m, was produced through a regression tree machine learning algorithm integrating GEDI LiDAR data and Landsat optical information. The second, with a spatial resolution of 10 m, was created using Sentinel-2 images and a deep learning convolutional neural network. We tested this approach separately in 30 forest plots situated in the northern Italian Alps, in 100 plots in the forested area of Traunstein (Germany) and successively in all the 130 plots through a cross-validation analysis. Forest density information was also included as influencing factor in a multiple regression analysis. Our results show that the GEDI CHMs can be used to assess biodiversity patterns in forest ecosystems through the estimation of the HH that is correlated to the tree species diversity. However, the results also indicate that this method is influenced by different factors including the GEDI CHMs dataset of choice and their related spatial resolution, the heterogeneity indices used to calculate the HH and the forest density. Our finding suggest that GEDI LIDAR data can be a valuable tool in the estimation of forest tree heterogeneity and related tree species diversity in forest ecosystems, which can aid in global biodiversity estimation.     

Temporal population variability in local forest communities has mixed effects on tree species richness across a latitudinal gradient

Among the local processes that determine species diversity in ecological communities, fluctuation‐dependent mechanisms that are mediated by temporal variability in the abundances of species populations have received significant attention. Higher temporal variability in the abundances of species populations can increase the strength of temporal niche partitioning but can also increase the risk of species extinctions, such that the net effect on species coexistence is not clear. We quantified this temporal population variability for tree species in 21 large forest plots and found much greater variability for higher latitude plots with fewer tree species. A fitted mechanistic model showed that among the forest plots, the net effect of temporal population variability on tree species coexistence was usually negative, but sometimes positive or negligible. Therefore, our results suggest that temporal variability in the abundances of species populations has no clear negative or positive contribution to the latitudinal gradient in tree species richness.     

Biodiversity assessment in incomplete inventories: leaf litter ant communities in several types of Bornean rain forest

Biodiversity assessment of tropical taxa is hampered by their tremendous richness, which leads to large numbers of singletons and incomplete inventories in survey studies. Species estimators can be used for assessment of alpha diversity, but calculation of beta diversity is hampered by pseudo-turnover of species in undersampled plots. To assess the impact of unseen species, we investigated different methods, including an unbiased estimator of Shannon beta diversity that was compared to biased calculations. We studied alpha and beta diversity of a diverse ground ant assemblage from the Southeast Asian island of Borneo in different types of tropical forest: diperocarp forest, alluvial forest, limestone forest and heath forests. Forests varied in plant composition, geology, flooding regimes and other environmental parameters. We tested whether forest types differed in species composition and if species turnover was a function of the distance between plots at different spatial scales. As pseudo-turnover may bias beta diversity we hypothesized a large effect of unseen species reducing beta diversity. We sampled 206 ant species (25% singletons) from ten subfamilies and 55 genera. Diversity partitioning among the four forest types revealed that whereas alpha species richness and alpha Shannon diversity were significantly smaller than expected, beta-diversity for both measurements was significantly higher than expected by chance. This result was confirmed when we used the unbiased estimation of Shannon diversity: while alpha diversity was much higher, beta diversity differed only slightly from biased calculations. Beta diversity as measured with the Chao-Sørensen or Morisita-Horn Index correlated with distance between transects and between sample points, indicating a distance decay of similarity between communities. We conclude that habitat heterogeneity has a high influence on ant diversity and species turnover in tropical sites and that unseen species may have only little impact on calculation of Shannon beta diversity when sampling effort has been high.     

Tree diversity on islands: assembly rules,passive sampling and the theory of island biogeography

Aim Species diversity is distributed heterogeneously through space, for reasons that are poorly understood. We tested three hypotheses to account for spatial variation in coniferous tree species diversity in a temperate island archipelago. The theory of island biogeography (ToIB) predicts that island area affects species diversity both directly (by increasing habitat diversity) and indirectly (by increasing abundances, which in turn reduce extinction rates). The ToIB also predicts that island isolation directly affects species diversity by reducing immigration rates. The passive sampling hypothesis predicts that island area and isolation both affect species diversity indirectly, by increasing and decreasing abundances, respectively. Community assembly rules (i.e. even partitioning of conifer abundances among islands) might also reduce tree species diversity beyond the core predictions of ToIB and the passive sampling hypothesis. Location Barkley Sound, British Columbia, Canada. Methods The abundances of eight coniferous tree species were quantified on 34 islands and two (1 ha) mainland plots. The predictions of the ToIB and the passive sampling hypothesis were tested using path analysis, and null models were used to test for abundance‐based assembly rules and to further test the passive sampling hypothesis. Results Path analysis showed that island area and isolation did not have direct, statistical effects on tree species diversity. Instead, both geographic variables had direct statistical effects on total tree abundances, which in turn predicted tree diversity. Results from several passive sampling null models were correlated with observed patterns in species diversity, but they consistently overestimated the number of tree species inhabiting most islands. A different suite of null models showed support for community assembly rules, or that tree species often reached higher abundances on islands that housed fewer heterospecific trees. Main conclusions Results were inconsistent with the ToIB. Instead, patterns in tree diversity were best explained by a combination of stochastic (passive sampling) and deterministic (assembly rules) processes. Stochastic and deterministic processes are commonly considered to be exclusive explanations for island community structure, but results from this study suggest that they can work synergistically to structure island tree communities.     

Small‐Sample Estimation of Species Richness Applied to Forest Communities

Summary Many well‐known methods are available for estimating the number of species in a forest community. However, most existing methods result in considerable negative bias in applications, where field surveys typically represent only a small fraction of sampled communities. This article develops a new method based on sampling with replacement to estimate species richness via the generalized jackknife procedure. The proposed estimator yields small bias and reasonably accurate interval estimation even with small samples. The performance of the proposed estimator is compared with several typical estimators via simulation study using two complete census datasets from Panama and Malaysia.     

Classification of vegetational diversity in managed boreal forests in eastern Finland

Diversity of vegetation in managed forests is studied. A classification based on forest stand structure, the abundances of vegetation species and variations in these abundances is developed and diversity indices are calculated for the classes to describe the diversity of the vegetation within the classes. The classes were formed using detrended correspondence analysis (DCA), global nonmetric multidimensional scaling (GNMDS) and TWINSPAN classification. Discriminant analysis was used to determine the environmental variables differentiating between the classes, and Duncan's multiple range test was used to examine the ability of the diversity measures to distinguish the classes. Beta diversity was estimated with Økland's method based on DCA ordination of the sample plots. The results point to fertility and the successional stage of the stand as the main factors affecting species diversity, in addition to which soil type, the number of tree species, crown cover, basal area and certain variables describing the management of the stand were relevant to the classification. The most distinct diversity indices were the reciprocal of Simpson, Pielou's J' and species richness, while the highest alpha diversity was found in young stands with a low crown cover on herb-rich or mesic forest sites. Beta diversity was quite high, its highest values of all for the whole data being recorded along the fertility gradient.     

西双版纳热带山地雨林群落乔木树种多样性研究

根据块样地资料对西双版纳热带山地雨林树种多样性特征进行了分析.结果表明,在2 500m²的样地上,随着起测胸径递增,各样地乔木个体数和树种丰富度均表现为依次递减;低海拔带上(850～1000 m)的山地雨林(1、2号样地)的各指数值无一定变化规律,较高海拔带上(1200～2000 m)的山地雨林(3～6号样地)的Shannon-Wiener指数和Simpson指数均表现为依次递减,而Pielou均匀度指数则呈依次递增的趋势.较高海拔带上的山地雨林在较小乔木起测胸径(≥2 cm、≥5 cm、≥10 cm)的树种丰富度、多样性和均匀度指数均要明显大于低海拔带上的山地雨林,而两者在较大乔木起测胸径(≥20 cm、≥30 cm、≥50 cm)的各指数值无明显差异.随着取样面积的递增,各样地树种丰富度、多样性和均匀度指数在取样面积递增到2 000m²处均已趋于平缓.     

National Forest Inventories in North America for monitoring forest tree species diversity

Abstract

A successful forest tree diversity-monitoring programme delivers reliable estimates of rates of occurrence, the spatial extent and the abundance of all forest-dwelling tree species. Sample-based estimators of these characteristics are provided for North American national forest inventories and discussed in the context of monitoring for forest tree diversity. The expected performance of the Canadian, the United States, and Mexican national forest inventory is quantified for three regions in each country. As expected, estimates for many less common and rare species are imprecise and sometimes these species are missed completely. We suggest augmenting existing national forest inventories by purposive sampling for these species.     

Efficient sampling of plant diversity in arid deserts using non-parametric estimators

Surveying plant diversity in arid desert areas is extremely difficult because of the harsh climate, hostile terrain, lack of roads, and insecurity, which is why it is particularly important to improve the sampling efficiency, but few relevant studies have been done. The performance of non-parametric estimators was assessed with first-hand field data to determine (a) the threshold of the proportion of uniques (number of species that occur in exactly one plot divided by the number of species sampled) that involves the least sampling effort and (b) the method of locating plots to obtain a more reliable estimate of species richness. The study area (Gurbantunggut desert, China) was divided into five sub-regions based on variation in physical environment and vegetation. The following common correction factors were selected: ACE, Chao1, Bootstrap, Chao2, ICE, Jack1, and Jack2. The estimates for each sub-region (partition) and for the entire region (without partition), the threshold of proportion of uniques, and the method of determining sampling locations (including prior sampling of plots that show large differences in habitats) were compared in terms of their ability to predict the number of species more accurately. We found that ACE and Chao1 (which use abundance data) showed more biased estimates than the other factors (incidence data), and best estimator is Jack1. Species richness was significantly underestimated for the region, but the non-parametric estimators could estimate the species richness for each sub-region reliably. Sampling locations affected the performance of non-parametric estimators significantly. The threshold of minimum sampling was 15% and that of uniques was 30%; the two were able to limit the bias within 5 and 10%, respectively. It is concluded that the non-parametric estimators can estimate the plant diversity of arid deserts reliably from the data on incidence. The study area (on the scale of a region) should be partitioned to improve the performance of the non-parametric estimators. The plots with larger differences in habitats should be sampled more extensively based on the threshold of the proportion of uniques.     

Up in the Tree – The Overlooked Richness of Bryophytes and Lichens in Tree Crowns

Assessing diversity is among the major tasks in ecology and conservation science. In ecological and conservation studies, epiphytic cryptogams are usually sampled up to accessible heights in forests. Thus, their diversity, especially of canopy specialists, likely is underestimated. If the proportion of those species differs among forest types, plot-based diversity assessments are biased and may result in misleading conservation recommendations. We sampled bryophytes and lichens in 30 forest plots of 20 m × 20 m in three German regions, considering all substrates, and including epiphytic litter fall. First, the sampling of epiphytic species was restricted to the lower 2 m of trees and shrubs. Then, on one representative tree per plot, we additionally recorded epiphytic species in the crown, using tree climbing techniques. Per tree, on average 54% of lichen and 20% of bryophyte species were overlooked if the crown was not been included. After sampling all substrates per plot, including the bark of all shrubs and trees, still 38% of the lichen and 4% of the bryophyte species were overlooked if the tree crown of the sampled tree was not included. The number of overlooked lichen species varied strongly among regions. Furthermore, the number of overlooked bryophyte and lichen species per plot was higher in European beech than in coniferous stands and increased with increasing diameter at breast height of the sampled tree. Thus, our results indicate a bias of comparative studies which might have led to misleading conservation recommendations of plot-based diversity assessments.     

Temporal Patterns of Species Accumulation in a Survey of Lepidoptera in a Beech-Maple Forest

One of the most significant challenges to insect conservation is lack of information concerning species diversity and distribution. Because a complete inventory of all species in an area is virtually impossible, interest has turned to developing statistical techniques to guide sampling design and to estimate total species richness within a site. We used two such techniques, diversity partitioning and non-parametric richness estimation, to determine how variation in sampling effort over time affected species accumulation for a survey of Lepidoptera in an old-growth beech-maple forest. Temporal scaling of sampling effort had significant effects on two measures of species diversity. Increases in species richness were primarily driven by changes in species occurrences with season, while Shannon diversity was largely determined at the scale of individual sampling units (i.e. by spatial effects). Variation in sampling effort affected the values of the two most widely regarded richness estimators (ICE and Chao 2); neither diversity estimator achieved stable values across a range of sampling efforts. Even after 52 trap-nights and accounting for seasonality, rare species (singletons and uniques) remained a significant component of the moth community. To the extent that moth communities in other forest systems are similarly comprised of many rare species, non-parametric richness estimators should be expected to yield variable estimates with increased effort and should only be used to provide a minimum benchmark for predicting the number of species remaining to be sampled. Our results suggest the best strategy for a short-term survey of forest Lepidoptera should emphasize spreading sampling intervals throughout a given year rather than focusing on intensive sampling during a short time period or prolonged sampling over many years.     

A novel method for estimating the number of species within a region

Ecologists are often required to estimate the number of species in a region or designated area. A number of diversity indices are available for this purpose and are based on sampling the area using quadrats or other means, and estimating the total number of species from these samples. In this paper, a novel theory and method for estimating the number of species is developed. The theory involves the use of the Laplace method for approximating asymptotic integrals. The method is shown to be successful by testing random simulated datasets. In addition, several real survey datasets are tested, including forests that contain a large number (tens to hundreds) of tree species, and an aquatic system with a large number of fish species. The method is shown to give accurate results, and in almost all cases found to be superior to existing tools for estimating diversity.     

Peatland forests are the least diverse tree communities documented in Amazonia,but contribute to high regional beta‐diversity

Western Amazonia is known to harbour some of Earth's most diverse forests, but previous floristic analyses have excluded peatland forests which are extensive in northern Peru and are among the most environmentally extreme ecosystems in the lowland tropics. Understanding patterns of tree species diversity in these ecosystems is important both for quantifying beta‐diversity in this region, and for understanding determinants of diversity more generally in tropical forests. Here we explore patterns of tree diversity and composition in two peatland forest types – palm swamps and peatland pole forests – using 26 forest plots distributed over a large area of northern Peru. We place our results in a regional context by making comparisons with three other major forest types: terra firme forests (29 plots), white‐sand forests (23 plots) and seasonally‐flooded forests (11 plots). Peatland forests had extremely low (within‐plot) alpha‐diversity compared with the other forest types that were sampled. In particular, peatland pole forests had the lowest levels of tree diversity yet recorded in Amazonia (20 species per 500 stems, Fisher's alpha 4.57). However, peatland pole forests and palm swamps were compositionally different from each other as well as from other forest types in the region. Few species appeared to be peatland endemics. Instead, peatland forests were largely characterised by a distinctive combination of generalist species and species previously thought to be specialists of other habitats, especially white‐sand forests. We suggest that the transient nature and extreme environmental conditions of Amazonian peatland ecosystems have shaped their current patterns of tree composition and diversity. Despite their low alpha‐diversity, the unique combination of species found in tree communities in Amazonian peatlands augment regional beta‐diversity. This contribution, alongside their extremely high carbon storage capacity and lack of protection at national level, strengthens their status as a conservation priority.     

Comparing tree diversity and composition in coffee farms and sacred forests in the Western Ghats of India

Expansion of coffee cultivation is one of the causes of deforestation and biodiversity loss. However, shade grown coffee has been promoted as a means for preserving biodiversity in the tropics. In this study we compared tree diversity in two types of coffee management regimes with the sacred groves in the Western Ghats of India. We computed species accumulation curves, species diversity indices and evenness indices to compare the different management regimes. Results of diversity indices showed that shade coffee had less diversity compared to sacred groves. Exotic species dominated the tree diversity in lands where the tree harvesting rights are with the growers. Native trees dominated the tree diversity when growers had no ownership rights on trees. A species accumulation curve suggested that the sacred grove had higher species richness compared to other two habitats. Lack of incentive to preserve endemic species as shade trees is forcing growers to plant more exotic species in shade grown coffee plots. If encouraged, shade grown coffee can preserve some biodiversity, but cannot provide all ecological benefits of a natural forest.     

Rapid Simultaneous Estimation of Aboveground Biomass and Tree Diversity Across Neotropical Forests: A Comparison of Field Inventory Methods

A standardized rapid inventory method providing information on both tree species diversity and aboveground carbon stocks in tropical forests will be an important tool for evaluating efforts to conserve biodiversity and to estimate the carbon emissions that result from deforestation and degradation (REDD). Herein, we contrast five common plot methods differing in shape, size, and effort requirements to estimate tree diversity and aboveground tree biomass (AGB). We simulated the methods across six Neotropical forest sites that represent a broad gradient in forest structure, tree species richness, and floristic composition, and we assessed the relative performance of methods by evaluating the bias and precision of their estimates of AGB and tree diversity. For a given sample of forest area, a ‘several small’ (< 1 ha) sampling strategy led to a smaller coefficient of variation (CV) in the estimate of AGB than a ‘few large’ one. The effort (person‐days) required to achieve an accurate AGB estimate (< 10% CV), however, was greater for the smallest plots (0.1 ha) than for a compromise approach using 0.5 ha modified Gentry plots, which proved to be the most efficient method to estimate AGB across all forest types. Gentry plots were also the most efficient at providing accurate estimates of tree diversity (< 10% CV of Hill number). We recommend the use of the 0.5 ha modified Gentry plot method in future rapid inventories, and we discuss a set of criteria that should inform any choice of inventory method.     

Species richness measures fail in resolving diversity patterns of speciose forest moth assemblages

We used data from a light-trapping study at 28 sites on floodplain forest moths in eastern Austria to assess the performance of a variety of species richness and species diversity measures. At each site the data (32,181 individuals from 448 species) contain a large fraction of species represented only as singletons. Sampling effort was evenly spread across sites, but sampling success varied greatly. Influx of moths from the landscape matrix surrounding floodplain forest patches lead to substantial proportions of stray individuals from the regional species pool. Under these conditions, observed species numbers as well as eight extrapolation estimators of species totals failed to reflect differences between three study regions or between flooded and non-flooded forest habitats. Rarefied species numbers and Fisher’s α of the log-series distribution captured differences in moth diversity between regions, but failed to mirror flooding impact. Only Shannon’s diversity captured all expected diversity differences, at high significance levels. Application of Chao and Shen’s bias correction increased figures of Shannon’s diversity, but did not affect the outcome of statistical comparisons. We conclude that for species-rich incompletely sampled communities of highly mobile insects the evaluation of the complete species-abundance information using Shannon’s diversity is the most promising mode to compare local species diversity with a high degree of ecological resolution. Species richness measures apart from those obtained through rarefaction cannot be recommended, as they are sensitive to sources of bias that pertain to many empirical sets of field data.     

Communities of ground-living spiders in deciduous forests: Does tree species diversity matter?

The relationships between species diversity and ecosystem functions are in the focus of recent ecological research. However, until now the influence of species diversity on ecosystem processes such as decomposition or mineral cycling is not well understood. In deciduous forests, spiders are an integral part of the forest floor food web. In the present study, patterns of spider diversity and community structure are related to diversity of deciduous forest stands in the Hainich National Park (Thuringia). In 2005, pitfall trapping and quantitative forest floor sampling were conducted in nine plots of forest stands with one (Diversity Level 1), three (DL 2) and five (DL 3) major deciduous tree species. Species richness, measured with both methods, as well as spider abundance in forest floor samples were highest in stands with medium diversity (DL 2) and lowest in pure beech stands (DL 1). The Shannon-Wiener index and spider numbers in pitfall traps decreased from DL 1 to DL 3, while the Shannon-Wiener index in forest floor samples increased in the opposite direction. Spider community composition differed more strongly between single plots than between diversity levels. Altogether, no general relationship between increasing tree species diversity and patterns of diversity and abundance in spider communities was found. It appears that there is a strong influence of single tree species dominating a forest stand and modifying structural habitat characteristics such as litter depth and herb cover which are important for ground-living spiders.     

A comparison of species diversity estimators

Although having been much criticized, diversity indices are still widely used in animal and plant ecology to evaluate, survey, and conserve ecosystems. It is possible to quantify biodiversity by using estimators for which statistical characteristics and performance are, as yet, poorly defined. In the present study, four of the most frequently used diversity indices were compared: the Shannon index, the Simpson index, the Camargo eveness index, and the Pielou regularity index. Comparisons were performed by simulating the Zipf–Mandelbrot parametric model and estimating three statistics of these indices, i.e., the relative bias, the coefficient of variation, and the relative root-mean-squared error. Analysis of variance was used to determine which of the factors contributed most to the observed variation in the four diversity estimators: abundance distribution model or sample size. The results have revealed that the Camargo eveness index tends to demonstrate a high bias and a large relative root-mean-squared error whereas the Simpson index is least biased and the Shannon index shows a smaller relative root-mean-squared error, regardless of the abundance distribution model used and even when sample size is small. Shannon and Pielou estimators are sensitive to changes in species abundance pattern and present a nonnegligible bias for small sample sizes (<1000 individuals). Received: May 8, 1998 / Accepted: May 6, 1999